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Three New Species of Pentagonia (Rubiaceae) from Southern Central America, One Foreseen, Two Surprising

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Three New Species of Pentagonia (Rubiaceae) from Southern Central America, One Foreseen, Two Surprising Hammel, B.E. 2015. Three new species of Pentagonia (Rubiaceae) from southern Central America, one foreseen, two surprising. Phytoneuron 2015-46: 1–13. Published 11 August 2015. ISSN 2153 733X THREE NEW SPECIES OF PENTAGONIA (RUBIACEAE) FROM SOUTHERN CENTRAL AMERICA, ONE FORESEEN, TWO SURPRISING BARRY E. HAMMEL Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166-0299 [email protected] ABSTRACT The species presented as "Pentagonia sp. A" in the recently published Vol. VII of Manual de Plantas de Costa Rica is here described as Pentagonia gomez-lauritoi Hammel, sp. nov. Two additional new species, also from Costa Rica, discovered and fully characterized too recently for inclusion in that volume, are described as well. The latter two are surprising not only for their untimely discovery but also for their unusual leaf morphology. Simple, entire leaves are the rule in Rubiaceae, with Pentagonia the one genus where species with pinnately lobed leaves have been described. Until now just 2 out of the 10 species known from Costa Rica (4 out of 15 in Flora Mesoamericana ) had lobed leaves; Pentagonia gambagam Hammel & Aguilar, sp. nov. , adds a third. The other new species, Pentagonia osapinnata Aguilar, N. Zamora, & Hammel, sp. nov. , was at the time of its discovery little short of astounding, as it has pinnately compound leaves, which had not been described for the family. RESUMEN La especie presentada como “ Pentagonia sp. A” en el recientemente publicado Vol. VII del Manual de Plantas de Costa Rica se describe aquí como P. gomez-lauritoi . Se describe dos especies adicionales, también de Costa Rica, descubiertas y caracterizadas tarde para incluirlas en dicho volumen. Estas dos últimas sorprenden, no solo por su reciente descubrimiento, sino también por la morfología inusual de sus hojas. Las hojas simples y enteras son la regla general en Rubiaceae, Pentagonia siendo el único género donde se ha descrito especies con hojas pinnadamente lobuladas. Hasta el momento se conocía solo dos especies (de las diez conocidas) con hojas lobuladas en Costa Rica (cuatro de 15 en Flora Mesoaméricana ); P. gambaga m figura como la tercera. La otra especie nueva, P. osapinnata , en el momento de su descubrimiento fue simplemente asomobrosa, ya que tiene hojas pinnadamente compuestas, lo cual no se había descrito para la familia. Pentagonia Benth. has not been revised in 100 years, during which time the number of recognized species has more than tripled; Standley (1914a) described five new species and recognized a total of eleven in his revision (as Watsonamra ). Including Nothophlebia costaricensis Standl., described in the same volume (Standley 1914b) and subsequently transferred to Pentagonia , a total of 12 species were known at the time. Including the three described here, approximately 40 species (12 in Costa Rica, 18 in Mesoamerica) are currently recognized and the genus is in dire need of a thorough revision with careful field work (Rova & Andersson 1995; Taylor 2012). The genus is easily recognized vegetatively by virtue of its frequently unbranched, subshrub to arborescent growth form with the leaves clustered towards the stem apex, its large, entire and strongly carinate, interpetiolar stipules, its usually rather large leaf blades (10 up to at least 200 cm long), and especially by the leaf surfaces distinctly and closely striate with fine epidermal fibers. The calyx tube usually has adaxial patches or lines of glands at or below the sinuses of the lobes. See also Rova and Andersson (1995) and Taylor (2002). It is the only genus of Rubiaceae with any species whose normal, mature leaves are pinnately lobed to deeply pinnatifid. In the New World, occasional individuals of some species of Simira may have pinnatifid leaves (Taylor 2002) and a few species of Cruckshanksia have leaves deeply and digitately 2 or 3 lobed (see Taylor 1996). Hammel: New Pentagonia , foreseen, surprising! 2 Contrary to what has sometimes been suggested elsewhere [cf. notes under Pentagonia donnell-smithii (Standl.) Standl. in Taylor 2012; Taylor & Hammel 2014] field work at La Selva research station and throughout Costa Rica demonstrates habit type to be quite constant within species of Pentagonia (see also Taylor 2002). Just three [ P. costaricensis (Standl.) W.C. Burger & C.M. Taylor, P. nuciformis Dwyer, and P. osaensis C.M. Taylor] of the 12 species now known from Costa Rica are typically much-branched trees up to 18 meters tall. All the others are treelets mostly less than 8 meters tall and usually monopodial. Of these latter, an occasional small plant may be somewhat branched from near the base, but otherwise in my experience any indicated habit outside these norms has been due to incorrect identification to species or a probable label mix-up. PENTAGONIA GAMBAGAM Hammel & Aguilar, sp. nov. TYPE : COSTA RICA. Puntarenas. Cantón de Golfito, Corredor Biológico la Gamba, Santuario Gambagam, 8°41' N, 83°11' W, 110 m, 28 Apr 2014 (fl, fr), B. Hammel, R. Aguilar , and I. Pérez 26730 (holotype: CR; isotypes: MO, GB). Figure 1. Similar to Pentagonia macrophylla by its reddish and bracteate inflorescence, its petiolate leaves, and its narrowly funnelform, trumpet-shaped corolla but differing markedly by its lobed leaf blades, more shallowly lobed calyx, and often greenish, totally glabrous corolla. It is also similar to the locally sympatric P. wendlandii , which has sessile or subsessile and entire leaves as well as a tubular and densely tomentulose corolla. The new species has leaves that are much more shallowly lobed than those of all other Pentagonia species in the region with lobed leaves. Shrub or small tree 4–8 m, unbranched; stems blackish brown, matte. Stipules reddish, 5–9 × 1.5–4.5 cm, ± ovate to deltate, acute to acuminate, coriaceous, abaxially and densely strigulose, adaxially glabrous. Leaves simple; petiole reddish green at the very base, to ca. 10 cm; blade 78–96 × 66–74 cm, spatulate to pandurate, basally truncate to more often shallowly cordate, rarely obtuse (often bullate between the basal secondary nerves), ± coriaceous, dull on both surfaces, densely and minutely strigulose on the main nerves abaxially, glabrous adaxially, pinnately lobed (1/3–2/3 the way to the midrib), and with 15–18 principal secondary veins per side (the ones feeding the lobes somewhat thicker than the rest), the lobes 4 or 5(–7) per side, 8–27 × 11–16 cm [the sinus ca. (4–)8– 12 cm from the midrib]. Inflorescences subsessile, glomerate; bracts reddish, to at least 30 × 16 mm, ovate, sparsely strigulose to glabrescent; flowers to at least 20, sessile or with pedicel to ca. 2 mm; hypanthium 5–7 mm, ± funnelform, densely strigulose; calyx limb reddish, 15–20 mm, coriaceous, basally and sparsely strigulose, apically glabrous, somewhat irregularly 5-lobed, the lobes imbricate, 5–7 mm, ± ovate to obovate, apically obtuse to rounded, the glands in lines 4–5 mm, from ca. mid tube downward; corolla green or greenish yellow to cream colored, 35–40[–55] mm, narrowly funnelform, trumpet-shaped, the tube basally 2–4[7] mm wide, apically 4–6[11] mm wide, straight to sometimes slightly curved at the base, internally pilosulose at the insertion of the filaments and pilosulose to strigulose upward to ca. 6 mm, otherwise glabrous inside and out, 5-lobed, the lobes 4–5[5–8] mm, triangular, marginally granulose puberulent except pilosulose at the very tip; stamens with the filament ca. 13 [15] mm, inserted at ca. 10 mm above the base of the tube, apically curved just below the anther (in life), basally pilosulose, the anthers ca. 3 mm, in life held perpendicular to the major portion of the filament and held at the same level or slightly above the stigma; style ca. 23 mm. Fruits reddish, 2.5–3.5 cm subglobose to ellipsoid, densely to sparsely strigulose and markedly pustulose-warty. Measurements in square brackets [ ] are from fresh or EtOH preserved material. Flowering Apr, Jun; fruiting Apr. At edge and in clearings of wet forest, 110 m elev.; Costa Rica (Prov. Puntarenas), Fila Gamba. Additional material examined. COSTA RICA. Puntarenas: Cantón de Golfito, Corredor Biológico la Gamba, Santuario Gambagam, 8°41' N, 83°11' W, 110 m, 2 Jun 2013 (fl), Aguilar 14395 (photos; see <https://www.flickr.com/search/?q=RA%2314395> ). Hammel: New Pentagonia , foreseen, surprising! 3 Etymology. This species is named (by a noun in apposition) for the Santuario Gambagam, its type and only known locality. Figure 1. Pentagonia gambagam . A. habit. B. Specimen before pressing, with leaf (showing lower surface), stem and inflorescence. C. Inflorescence; D. Flower showing corolla split (on each side of the flower from which removed), flower bud, and associated bract. E. fruit split open. Scale bars: B=20 cm, C=4 cm, D & E=2 cm. All from Hammel et al. 26730 . Hammel: New Pentagonia , foreseen, surprising! 4 Figure 2. Pentagonia macrophylla . Inflorescence from Hammel et al. 26294 (MO). Figure 3. Pentagonia wendlandii. Inflorescence from Hammel et al. 26733 (CR). Hammel: New Pentagonia , foreseen, surprising! 5 Figure 4. Pentagonia lobata . Inflorescence from Hammel et al. 26114 (CR). Because of its monopodial habit, large, petiolate leaves, reddish inflorescence with reddish, relatively large bracts, and funnelform, trumpet-shaped flowers, this species on first impression seemed to be an odd version of Pentagonia macrophylla Benth. with lobed leaves. However, closer examination prevented us from leaving it lie as that. In any case, the taxonomy we follow here, that of Taylor 1995, 2002, 2012 and Taylor & Hammel 2014) gives considerable weight to lobate vs. entire leaves as an indicator of speciation. Armed with that consideration, we find that the more shallowly lobed calyx (ca. 1/3 vs. 3/4 to completely), the often greenish and externally glabrous (vs. white to cream-colored and densely strigose-tomentulose) corolla along with the lobate leaves clearly distinguish P.
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